Chapter 44

Osmoregulation and Excretion

PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero

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 Overview: A balancing act The physiological systems of animals

Operate in a fluid environment

The relative concentrations of water and solutes in this environment

Must be maintained within fairly narrow limits

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 Freshwater animals
Show adaptations that reduce water uptake and conserve solutes

Desert and marine animals face desiccating environments

With the potential to quickly deplete the body water

Figure 44.1
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 Osmoregulation
Regulates solute concentrations and balances the gain and loss of water

Excretion
Gets rid of metabolic wastes

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 Concept 44.1: Osmoregulation balances the uptake and loss of water and solutes Osmoregulation is based largely on controlled movement of solutes
Between internal fluids and the external environment

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Osmosis Cells require a balance

Between osmotic gain and loss of water

Water uptake and loss

Are balanced by various mechanisms of osmoregulation in different environments

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Osmotic Challenges Osmoconformers, which are only marine animals

Are isoosmotic with their surroundings and do not regulate their osmolarity

Osmoregulators expend energy to control water uptake and loss

In a hyperosmotic or hypoosmotic environment

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 Most animals are said to be stenohaline

And cannot tolerate substantial changes in external osmolarity

Euryhaline animals
Can survive large fluctuations in external osmolarity

Figure 44.2
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Marine Animals Most marine invertebrates are osmoconformers Most marine vertebrates and some invertebrates are osmoregulators

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 Marine bony fishes are hypoosmotic to sea water

And lose water by osmosis and gain salt by both diffusion and from food they eat

These fishes balance water loss

By drinking seawater
Gain of water and salt ions from food and by drinking seawater Osmotic water loss through gills and other parts of body surface

Excretion of salt ions from gills

Excretion of salt ions and small amounts of water in scanty urine from kidneys

Figure 44.3a
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(a) Osmoregulation in a saltwater fish

Freshwater Animals Freshwater animals

Constantly take in water from their hypoosmotic environment Lose salts by diffusion

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 Freshwater animals maintain water balance

By excreting large amounts of dilute urine

Salts lost by diffusion

Are replaced by foods and uptake across the gills
Osmotic water gain through gills and other parts of body surface Uptake of water and some ions in food

Uptake of salt ions by gills

Excretion of large amounts of water in dilute urine from kidneys

Figure 44.3b (b) Osmoregulation in a freshwater fish

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Animals That Live in Temporary Waters Some aquatic invertebrates living in temporary ponds
Can lose almost all their body water and survive in a dormant state

This adaptation is called anhydrobiosis

100 m

100 m

Figure 44.4a, b

(a) Hydrated tardigrade

(b) Dehydrated tardigrade

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Land Animals Land animals manage their water budgets

By drinking and eating moist foods and by using metabolic water
Water balance in a kangaroo rat (2 mL/day = 100%) Ingested in food (0.2) Water gain Ingested in liquid (1,500) Water balance in a human (2,500 mL/day = 100%)

Ingested in food (750)

Derived from metabolism (1.8)

Derived from metabolism (250)

Feces (0.9)
Urine (0.45) Urine (1,500)

Feces (100)

Water loss

Figure 44.5
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Evaporation (1.46)

Evaporation (900)

 Desert animals
Get major water savings from simple anatomical features
EXPERIMENT Knut and Bodil Schmidt-Nielsen and their colleagues from Duke University observed that the fur of camels exposed to full sun in the Sahara Desert could reach temperatures of over 70C, while the animals skin remained more than 30C cooler. The Schmidt-Nielsens reasoned that insulation of the skin by fur may substantially reduce the need for evaporative cooling by sweating. To test this hypothesis, they compared the water loss rates of unclipped and clipped camels. RESULTS Removing the fur of a camel increased the rate of water loss through sweating by up to 50%.

Water lost per day (L/100 kg body mass)

4 3 2 1 0 Control group (Unclipped fur) Experimental group (Clipped fur)

CONCLUSION

The fur of camels plays a critical role in their conserving water in the hot desert environments where they live.

Figure 44.6
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Transport Epithelia Transport epithelia

Are specialized cells that regulate solute movement Are essential components of osmotic regulation and metabolic waste disposal

Are arranged into complex tubular networks

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 An example of transport epithelia is found in the salt glands of marine birds

Which remove excess sodium chloride from the blood
Nasal salt gland (a) An albatrosss salt glands empty via a duct into the nostrils, and the salty solution either drips off the tip of the beak or is exhaled in a fine mist. Nostril with salt secretions

Lumen of secretory tubule Vein Capillary Secretory tubule Transport epithelium Direction of salt movement Artery
NaCl

(b) One of several thousand secretory tubules in a saltexcreting gland. Each tubule is lined by a transport epithelium surrounded by capillaries, and drains into a central duct.

Blood Secretory cell flow of transport epithelium Central duct

(c) The secretory cells actively transport salt from the blood into the tubules. Blood flows counter to the flow of salt secretion. By maintaining a concentration gradient of salt in the tubule (aqua), this countercurrent system enhances salt transfer from the blood to the lumen of the tubule.

Figure 44.7a, b

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 Concept 44.2: An animals nitrogenous wastes reflect its phylogeny and habitat The type and quantity of an animals waste products
May have a large impact on its water balance

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 Among the most important wastes

Are the nitrogenous breakdown products of proteins and nucleic acids
Proteins

Nucleic acids

Amino acids

Nitrogenous bases

NH2 Amino groups

Many reptiles Most aquatic Mammals, most (including animals, including amphibians, sharks, birds), insects, most bony fishes some bony fishes land snails O C O C NH2 NH2 Urea O HN C H N C C O

NH3

C N N H H Uric acid

Figure 44.8

Ammonia

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Forms of Nitrogenous Wastes Different animals

Excrete nitrogenous wastes in different forms

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Ammonia Animals that excrete nitrogenous wastes as ammonia

Need access to lots of water Release it across the whole body surface or through the gills

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Urea The liver of mammals and most adult amphibians

Converts ammonia to less toxic urea

Urea is carried to the kidneys, concentrated

And excreted with a minimal loss of water

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Uric Acid Insects, land snails, and many reptiles, including birds
Excrete uric acid as their major nitrogenous waste

Uric acid is largely insoluble in water

And can be secreted as a paste with little water loss

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The Influence of Evolution and Environment on Nitrogenous Wastes The kinds of nitrogenous wastes excreted
Depend on an animals evolutionary history and habitat

The amount of nitrogenous waste produced

Is coupled to the animals energy budget

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 Concept 44.3: Diverse excretory systems are variations on a tubular theme Excretory systems
Regulate solute movement between internal fluids and the external environment

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Excretory Processes Most excretory systems

Produce urine by refining a filtrate derived from body fluids
Capillary Excretory tubule 1 Filtration. The excretory tubule collects a filtrate from the blood. Water and solutes are forced by blood pressure across the selectively permeable membranes of a cluster of capillaries and into the excretory tubule.

Filtrate
Urine

2 Reabsorption. The transport epithelium reclaims valuable substances from the filtrate and returns them to the body fluids.

3 Secretion. Other substances, such as toxins and excess ions, are extracted from body fluids and added to the contents of the excretory tubule.

4 Excretion. The filtrate leaves the system and the body.

Figure 44.9
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 Key functions of most excretory systems are

Filtration, pressure-filtering of body fluids producing a filtrate Reabsorption, reclaiming valuable solutes from the filtrate

Secretion, addition of toxins and other solutes from the body fluids to the filtrate
Excretion, the filtrate leaves the system

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Survey of Excretory Systems The systems that perform basic excretory functions
Vary widely among animal groups Are generally built on a complex network of tubules

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Protonephridia: Flame-Bulb Systems A protonephridium

Is a network of dead-end tubules lacking internal openings
Nucleus of cap cell Cilia Interstitial fluid filters through membrane where cap cell and tubule cell interdigitate (interlock) Tubule cell

Flame bulb Protonephridia (tubules)

Tubule

Figure 44.10
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Nephridiopore in body wall

 The tubules branch throughout the body

And the smallest branches are capped by a cellular unit called a flame bulb

These tubules excrete a dilute fluid

And function in osmoregulation

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Metanephridia Each segment of an earthworm

Has a pair of open-ended metanephridia

Coelom Capillary network Bladder Collecting tubule

Nephridiopore

Figure 44.11

Nephrostome

Metanephridia

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 Metanephridia consist of tubules

That collect coelomic fluid and produce dilute urine for excretion

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Malpighian Tubules In insects and other terrestrial arthropods, malpighian tubules

Remove nitrogenous wastes from hemolymph and function in osmoregulation
Digestive tract Rectum Intestine Hindgut

Malpighian Midgut tubules (stomach)

Salt, water, and nitrogenous wastes

Feces and urine

Anus

Malpighian tubule Rectum

Reabsorption of H2O, ions, and valuable organic molecules

Figure 44.12

HEMOLYMPH

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 Insects produce a relatively dry waste matter

An important adaptation to terrestrial life

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Vertebrate Kidneys Kidneys, the excretory organs of vertebrates

Function in both excretion and osmoregulation

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 Concept 44.4: Nephrons and associated blood vessels are the functional unit of the mammalian kidney The mammalian excretory system centers on paired kidneys
Which are also the principal site of water balance and salt regulation

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 Each kidney
Is supplied with blood by a renal artery and drained by a renal vein

Posterior vena cava

Renal artery and vein Aorta Ureter Urinary bladder Urethra (a) Excretory organs and major associated blood vessels Kidney

Figure 44.13a

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 Urine exits each kidney

Through a duct called the ureter

Both ureters
Drain into a common urinary bladder

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Structure and Function of the Nephron and Associated Structures

The mammalian kidney has two distinct regions

An outer renal cortex and an inner renal medulla
Renal medulla Renal cortex Renal pelvis

Ureter Section of kidney from a rat

Figure 44.13b

(b) Kidney structure

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 The nephron, the functional unit of the vertebrate kidney

Consists of a single long tubule and a ball of capillaries called the glomerulus
JuxtaCortical medullary nephron nephron
Afferent arteriole from renal artery

Glomerulus
Bowmans capsule Proximal tubule Peritubular capillaries

Renal cortex

Collecting duct

20 m Renal medulla

SEM Efferent arteriole from glomerulus Branch of renal vein Descending limb Ascending limb Vasa recta Distal tubule Collecting duct

To renal pelvis

Loop of Henle

Figure 44.13c, d
(c) Nephron Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings (d) Filtrate and blood flow

Filtration of the Blood Filtration occurs as blood pressure

Forces fluid from the blood in the glomerulus into the lumen of Bowmans capsule

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 Filtration of small molecules is nonselective

And the filtrate in Bowmans capsule is a mixture that mirrors the concentration of various solutes in the blood plasma

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Pathway of the Filtrate From Bowmans capsule, the filtrate passes through three regions of the nephron
The proximal tubule, the loop of Henle, and the distal tubule

Fluid from several nephrons

Flows into a collecting duct

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Blood Vessels Associated with the Nephrons

Each nephron is supplied with blood by an afferent arteriole
A branch of the renal artery that subdivides into the capillaries

The capillaries converge as they leave the glomerulus

Forming an efferent arteriole

The vessels subdivide again

Forming the peritubular capillaries, which surround the proximal and distal tubules

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From Blood Filtrate to Urine: A Closer Look Filtrate becomes urine

As it flows through the mammalian nephron and collecting duct
1 Proximal tubule NaCl Nutrients HCO3 H2O K+ 4 Distal tubule NaCl H2O HCO3

H+

NH3

K+

H+

CORTEX Filtrate H2O Salts (NaCl and others) HCO3 H+ Urea Glucose; amino acids Some drugs 2 Descending limb of loop of Henle H2O OUTER MEDULLA 3 Thin segment of ascending limb NaCl 5 Collecting duct

3 Thick segment of ascending limb

NaCl

Key Active transport Passive transport NaCl INNER MEDULLA

Urea
H2O

Figure 44.14
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 Secretion and reabsorption in the proximal tubule

Substantially alter the volume and composition of filtrate

Reabsorption of water continues

As the filtrate moves into the descending limb of the loop of Henle

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 As filtrate travels through the ascending limb of the loop of Henle

Salt diffuses out of the permeable tubule into the interstitial fluid

The distal tubule

Plays a key role in regulating the K+ and NaCl concentration of body fluids

The collecting duct

Carries the filtrate through the medulla to the renal pelvis and reabsorbs NaCl

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 Concept 44.5: The mammalian kidneys ability to conserve water is a key terrestrial adaptation The mammalian kidney
Can produce urine much more concentrated than body fluids, thus conserving water

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Solute Gradients and Water Conservation In a mammalian kidney, the cooperative action and precise arrangement of the loops of Henle and the collecting ducts
Are largely responsible for the osmotic gradient that concentrates the urine

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 Two solutes, NaCl and urea, contribute to the osmolarity of the interstitial fluid
Which causes the reabsorption of water in the kidney and concentrates the urine
300 300 300 100 300 CORTEX Active transport H2O H2O H2O H2O H2O H2O INNER MEDULLA H2O 400 NaCl 200 H2O 400 300 100

Osmolarity of interstitial fluid (mosm/L)

NaCl NaCl NaCl

Passive transport

H2O H2O H2O H2O Urea

400

OUTER MEDULLA

600
NaCl NaCl NaCl

400

600

600

900

700

H2O Urea H2O Urea 1200

900

1200

1200

Figure 44.15
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 The countercurrent multiplier system involving the loop of Henle

Maintains a high salt concentration in the interior of the kidney, which enables the kidney to form concentrated urine

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 The collecting duct, permeable to water but not salt

Conducts the filtrate through the kidneys osmolarity gradient, and more water exits the filtrate by osmosis

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 Urea diffuses out of the collecting duct

As it traverses the inner medulla

Urea and NaCl

Form the osmotic gradient that enables the kidney to produce urine that is hyperosmotic to the blood

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Regulation of Kidney Function The osmolarity of the urine

Is regulated by nervous and hormonal control of water and salt reabsorption in the kidneys

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 Antidiuretic hormone (ADH)

Increases water reabsorption in the distal tubules and collecting ducts of the kidney
Osmoreceptors in hypothalamus Hypothalamus Drinking reduces blood osmolarity to set point ADH Increased permeability Pituitary gland Distal tubule Thirst

STIMULUS: The release of ADH is triggered when osmoreceptor cells in the hypothalamus detect an increase in the osmolarity of the blood

H2O reabsorption helps prevent further osmolarity increase Collecting duct

Homeostasis: Blood osmolarity

Figure 44.16a

(a) Antidiuretic hormone (ADH) enhances fluid retention by making the kidneys reclaim more water.

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 The renin-angiotensin-aldosterone system (RAAS)

Is part of a complex feedback circuit that functions in homeostasis
Homeostasis: Blood pressure, volume Increased Na+ and H2O reabsorption in distal tubules STIMULUS: The juxtaglomerular apparatus (JGA) responds to low blood volume or blood pressure (such as due to dehydration or loss of blood) Arteriole constriction Adrenal gland

Aldosterone

Angiotensin II Distal tubule Angiotensinogen JGA

Renin production Renin

Figure 44.16b

(b) The renin-angiotensin-aldosterone system (RAAS) leads to an increase in blood volume and pressure.

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 Another hormone, atrial natriuretic factor (ANF)

Opposes the RAAS

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 The South American vampire bat, which feeds on blood

Has a unique excretory system in which its kidneys offload much of the water absorbed from a meal by excreting large amounts of dilute urine

Figure 44.17
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 Concept 44.6: Diverse adaptations of the vertebrate kidney have evolved in different environments The form and function of nephrons in various vertebrate classes
Are related primarily to the requirements for osmoregulation in the animals habitat

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 Exploring environmental adaptations of the vertebrate kidney

MAMMALS
BIRDS AND OTHER REPTILES

Bannertail Kangaroo rat (Dipodomys spectabilis)

Roadrunner (Geococcyx californianus)

Beaver (Castor canadensis) FRESHWATER FISHES AND AMPHIBIANS

Desert iguana (Dipsosaurus dorsalis) MARINE BONY FISHES

Rainbow trout (Oncorrhynchus mykiss)

Northern bluefin tuna (Thunnus thynnus)

Figure 44.18

Frog (Rana temporaria)

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